Genet Resour Crop Evol

https://doi.org/10.1007/s10722-018-0659-9

NOTES ON NEGLECTED AND UNDERUTILIZED CROPS

Vegetative propagation of the underutilized oilseed crop

sacha inchi (Plukenetia volubilis L.)
Danter Huansi Cachique . Henry Ruiz Solsol . Marco Antonio Garcı́a Sanchez .
Luis Alberto Arévalo López . Nete Kodahl

Received: 16 October 2017 / Accepted: 12 June 2018

Ó Springer Nature B.V. 2018

Abstract Underutilized crop species have the poten-
tial to aid in alleviating some of the challenges faced
by the world today; reduced food security and
malnutrition, land degradation, and climate change.
The incorporation of little-known crops into sustain-
able agricultural systems provides new developmental
opportunities and can increase the resilience of food
production systems. However, measures need to be
taken to address issues of variation in product quality
and access to propagation material. Plukenetia volu-
bilis L. (Euphorbiaceae) is an oleaginous liana native
to the Amazon basin. It is of growing economic
importance in the food, pharmaceutic, cosmetic and
horticultural industries, however, commercial culti-
vars are lacking and the genetic variation within the
species is very high. Furthermore, the majority of
landraces of P. volubilis are susceptible to nematodes.
In order to attain plants with a consistent quality and
yield, vegetative propagation schemes are beneficial.
D. H. Cachique
M. A. G. Sanchez
L. A. A. López
Instituto de Investigaciones de la Amazonia Peruana
(IIAP), Av. José A. Quiñones km 2.5, Iquitos, Peru
Grafting is an especially favourable technique which
enables the selection of pest and disease resistant
rootstocks in combination with high-yielding or
ornamentally valuable scions. Three different grafting
techniques combined with three different systems of
protection were tested on P. volubilis and evaluated
according to the growth of the scions following
grafting. The experiment was carried out under
nursery conditions in the San Martı́n region of Peru,
where P. volubilis is commonly cultivated. Regardless
of the grafting technique, a 100% survival rate was
found for plants covered with a plastic bag following
grafting. This result may aid in the further domesti-
cation of P. volubilis by providing farmers with access
to plant material of consistent quality and allowing the
selection of pest resistant rootstocks in combination
with high-yielding scions.
Keywords Rootstocks
Disease resistance
Genetic
homogeneity
Inca peanut
Grafting techniques

Underutilized plants

H. R. Solsol
Centro Agronómico Tropical de Investigación y
Introduction
Enseñanza (CATIE), Cartago, Costa Rica
The genus Plukenetia L. (Euphorbiaceae) includes 21
N. Kodahl (&) species of lianas and scrambling vines, of which 14 are
Department of Plant and Environmental Sciences,
University of Copenhagen, Thorvaldsensvej 40,
found in the Neotropics (Gillespie 2007; Bussmann
1871 Frederiksberg, Denmark et al. 2009, 2013). The most commonly cultivated
e-mail: nk@plen.ku.dk

123

species is Plukenetia volubilis L., which is distributed
across the Lesser Antilles, Surinam, and along the
Northern and Western edge of the Amazon basin in
Venezuela, Colombia, Ecuador, Peru, Bolivia, and
Brazil (Gillespie 1993). Plukenetia volubilis is a
monoecious, herbaceous to woody perennial climber
with ovate to cordate leaves, and several staminate
flowers situated above one to few pistillate flowers in a
racemose thyrse. The flowers are small, with greenish
to cream sepals and no tepals, and the fruits are large,
carinate to winged capsules with four to six carpels
and lenticular seeds (Gillespie 1993). The seeds
contain ca. 35–60% oil and have traditionally been
collected and consumed by the indigenous peoples of
the Amazon region (Chirinos et al. 2013; Gutiérrez
et al. 2011; Hamaker et al. 1992; Jáuregui et al. 2013;
Ruiz-Solsol and Mesén 2010). The native peoples of
Peru consume oil and flour produced from the seeds,
both for alimentation, and as a traditional medicine,
especially for controlling cholesterol levels (Aspajo
et al. 2016). In Ecuador, the seeds may be crushed with
cocoa (Theobroma cacao L.) for use in a hot beverage
(pers. comm. Ovidio Bolaños, June 2016). It has also
been cultivated in the Antilles, where the leaves are
eaten as vegetables or used as fodder (Hanelt 2001).
In recent years, P. volubilis has gained increasing
popularity in the food, pharmaceutical, cosmetic
and—most recently—horticultural industries, under
various local names such as sacha inchi (forest
peanut), Inca peanut, mani de estrella (star peanut)
and Inca nut. The seed oil includes approximately 45%
linolenic (omega-3) and 35% linoleic (omega-6) acids,
with oleic, palmitic and stearic fatty acids only present
in smaller amounts (Gutiérrez et al. 2011; Hamaker
et al. 1992). The polyunsaturated fatty acids omega-3
and -6 are known to have a positive influence on
human health, e.g., by preventing cardiovascular
diseases, including coronary heart disease and hyper-
tension (Cisneros et al. 2014; Fanali et al. 2011;
Guillén et al. 2003) making P. volubilis oil highly
attractive for the food and pharmaceutical industries.
Furthermore, the seeds contain antioxidants, including
vitamins A and E, (Chirinos et al. 2013) which may
explain the oil’s increasing popularity in the cosmetic
industry. The nuts are commonly consumed roasted
and salted, but may also be sold as flour or included in
granola, and the seed cake can be found marketed as
‘protein powder’.
123
Genet Resour Crop Evol
A novel trend is the use of P. volubilis as an
ornamental. In cut flower arrangements the large, star
shaped fruits in combination with the grass green
colour of the ovate to cordate leaves are very attractive
(pers. comm. César Cachique Huansi, September
2017). Furthermore, although the flowers are incon-
spicuous, the male flowers are elegant and fit well with
the trend of flower arrangements with a ‘‘wild’’ look.
In Ecuador, it is also common to include a single plant
of P. volubilis in home gardens, partly for culinary
purposes, but also for the aesthetic value of the vine
and the fruits (pers. comm. Ovidio Bolaños, June
2016).
Plukenetia volubilis has a large commercial poten-
tial, but so far breeding approaches and genetic
improvement have been extremely limited (Kri-
vankova et al. 2012). There is a considerable degree
of morphological variation between populations
(Arévalo 1996; Krivankova et al. 2012) and a high
genetic diversity between different populations (Co-
razon-Guivin et al. 2008, 2009). This effect increases
with the geographic distances between populations
(Rodrigues et al. 2013). Vegetative propagation tech-
niques allow for the conservation of genotypes and
maintenance of germplasm collections, by enabling
continued survival of selected genotypes (Zobel and
Talbert 1988). Furthermore, in the absence of com-
mercial cultivars, vegetative propagation techniques
are useful for rapidly multiplying superior genotypes
(Barter 2016; Zobel and Talbert 1988), which is
especially beneficial for allogamous species with
heterogeneous offspring such as P. volubilis (Cachi-
que 2006). Multiplication of high quality plants can
infer benefits to the local people in the plant’s native
range, e.g., through sustainable income generation and
by shifting cultivation from illicit crops such as
Erythroxylum coca Lam.
Vegetative propagation of P. volubilis through
cuttings has previously been demonstrated by Ruiz-
Solsol and Mesén (2010) and Cachique et al. (2011)
using indole-3-butyric acid for root induction. How-
ever, vegetative propagation of P. volubilis through
grafting has to our knowledge not previously been
attempted. Grafting is beneficial since it allows for the
selection of rootstocks and scions with different traits,
e.g., disease resistance and high yield, respectively
(Barter 2016). Furthermore, grafting with mature
scions can yield plants that will produce fruit earlier
than if the plant was propagated by seeds (Barter
Genet Resour Crop Evol
2016), thus reducing the time from the establishment
of a new plantation to the first harvest.
Grafting requires severing of the vascular system of
both the rootstock and scion and stops water and
nutrient transport throughout the plant. Successful
grafting therefore depends on restoration of the
vascular system through the division of cells in the
graft union (Parkinson et al. 1990). This requires that
the wound is not infected by pathogens, e.g., fungi, and
that the graft union is not subject to desiccation. While
grafting is most commonly used for woody species,
herbaceous plants can also be grafted (Parkinson et al.
1990).
The current study aims to analyse whether it is
possible to graft P. volubilis and, if so, to ascertain
which grafting method and protection system are the
most efficient.
Materials and methods
Environmental conditions
The present experiment was carried out at the
experimental nursery at Instituto de Investigaciones
de la Amazonı́a Peruana (IIAP, the Peruvian Amazon
Research Institute). IIAP is situated at an altitude of
332 m a.s.l., has an average temperature and monthly
precipitation of 22.61 °C and 70.02 mm, respectively,
and an average relative humidity of 77.80% (Estación
Metereológica del INIA, El porvenir, 2016).
Plant material
Rootstocks
Plukenetia volubilis L. (Fig. 1a) was used as rootstock
and the plants were produced at the nursery at IIAP
2 months prior to the experiment. Seeds for cultivation
of the rootstock were collected from Centro de
Investigaciones ‘‘Pucayacu’’ del IIAP and were chosen
because the mother plant has shown resistance to
nematodes and fungi compared to other plants in the
plantation. The rootstocks were grown from seeds in
black seedling bags of ca. 15 9 30 cm, with watering
every second day, regular application of foliar NPK
fertilizer 35-6-10 at 4.5 g/L and inhibition of fungal
growth through the application of
metalaxil ? mancozeb 2 g/L. At the time of the
experiment the plants were approximately 80 cm tall.
Prior to grafting, the lowest leaves were removed
and the rootstock plants were cut down to 12.66 cm in
height (Fig. 1b), in order for the diameter at the stem
section to be at 0.52 cm, coinciding with that of the
scion. The whole process was carried out beneath a
shade with 20% light transparency.
Scions
Plukenetia volubilis was also used as a scion, and the
plant material was taken from 1‘ year old plants from
the collection grown at the Centro de Investigaciones
‘‘Pucayacu’’ del IIAP. The collection comprises plants
grown from seeds collected in the San Martı́n region of
Peru. This accession was selected based on desirable
phenotypical characteristics; the plants have a high
yield of seeds based on weight and the seeds are large.
In order to optimize plant health prior to grafting,
the plants were prepared 1 month before harvest of the
grafting material. This was done through application
of bencimidazole 1 mL/L to reduce fungal growth,
foliar NPK fertilizer 35–6–10 at 4.5 g/L and soil NPK
fertilizer 120–70–90 to improve plant vigour.
The most vigorous shoots were selected as scions
and were collected between 5:30 and 7:30 a.m. in the
morning. The leaves were removed from the scions,
while care was taken to avoid damaging the axillary
buds. Shoots 30–40 cm in length were excised and
were transferred to the propagation area in coolers,
placed between layers of damp paper and labelled with
the origin of the plant.
Grafting procedure and experimental design
The grafting was performed under a cover with 20%
light transparency, and under aseptic conditions to
avoid tissue dehydration and contamination. Using
pruning shears disinfected in 96% alcohol, 1 cm stem
pieces were removed from both ends of the 40 cm long
shoots that were to be used for scions, in order to
eliminate any air bubbles that might have formed.
Subsequently, each of the shoots were divided into 2–3
scions of 10–15 cm in length, each scion containing
3–4 axillary buds with shooting potential.
The grafting experiment was set up in a randomized
block design. Each block contained 3 9 3 fields
consisting of 9 different treatments, and the blocks
123

Fig. 1 Grafting of Plukenetia volubilis; a mature plant with
fruit and unisexual flowers, b the rootstock is cut to 12.66 cm in
height, c plant prepared for straight grafting, d cleft grafting,
were replicated four times. Each of the 9 fields in the
blocks contained 10 grafted plants of P. volubilis,
including a total of 360 plants in the study. The nine
different treatments consisted of three different graft-
ing methods; cleft, straight and splice (see Figs. 1c, d
and 2a, b, c) and three different protection schemes;
sealing the wound with a plastic strip (Fig. 1e) and
covering the plant with a plastic bag (Fig. 1f), or
sealing the wound with a plastic strip and leaving the
plant uncovered, or sealing the wound with liquid
paraffin and leaving the plant uncovered.
123
Genet Resour Crop Evol
e sealing of grafting wound with plastic strip, f protection of
grafted plant with plastic bag
The grafted plants were placed under a shade
allowing 20% of the natural light to pass through. The
environmental conditions under the shade were mea-
sured during the first 5 days after grafting. The indoor
relative humidity varied between 66 and 89%, the
average temperature was 27 °C and the average solar
radiation was 75 lux.
All measurements were performed manually
45 days after grafting; % scion survival rate, %
mortality, number of shoots, length of shoots and
diameter of shoots.
Genet Resour Crop Evol
Fig. 2 Schematic
illustration of the three
grafting techniques;
a straight cut b cleft c splice
Data analysis
The percentages were arcsine transformed and the
counting data were transformed by squareroot(x ? 1)
(Snedecor and Cochran 1980). An analysis of variance
(ANOVA) test was used to determine significant
differences between grafting techniques and protec-
tion schemes, as well as differences arising through the
interactions between grafting techniques and protec-
tion schemes. Statistical differences between the
different grafting techniques were determined using
a Duncan’s new multiple range test (MRT), and the
differences between the protection schemes were
determined similarly.
Results
There were no significant differences in the percentage
of graft unions formed, percentage of mortality,
number of shoots, length of shoots, and diameter of
shoots between the cleft, straight cut and splice
grafting techniques (p [ 0.05). However, while the
graft union success was similar for the cleft and
straight cut techniques (56.67%), it was slightly lower
for the splice technique (51.67%, Table 1).
The protection systems yielded significantly differ-
ent results for both the percentage of graft unions
formed, the percentage of mortality, and the number,
length, and diameter of shoots (p \ 0.05, Table 2).
Protection of the grafted plants with a plastic bag
resulted in 100% graft union success and 0% mortal-
ity, which was significantly different from the 42.5 and
22.5% graft union success of the plants with no
123
 Genet Resour Crop Evol

Table 1 Effects of the grafting techniques on the growth of the grafted P. volubilis evaluated 45 days after grafting
Grafting technique Graft unions formed (%) Mortality (%) Shoots (No.) Length of shoots (cm) Diameter of shoots
(cm)

Cleft 56.67a 40.83a 1.78a 5.51a 0.22a

a a a a
Straight 56.67 40.00 1.71 5.27 0.21a
a a a a
Splice 51.67 45.83 1.77 5.36 0.24a
C.V. (%) 3.27 5.66 1.31 4.63 1.36
Average 55.00 42.22 1.75 5.38 0.22
Average values in each column followed by the same letter are statistically equal (Duncan’s MRT, p [ 0.05). C.V, coefficient of
variation

Table 2 Effects of the protection systems on the growth of the grafted P. volubilis evaluated 45 days after grafting
Protection system Graft unions formed (%) Mortality (%) Shoots (No.) Length of shoots (cm) Diameter of shoots
(cm)

Plastic bag 100.00a 0.00c 2.23a 5.51a 0.22b

c a c c
Liquid paraffin 22.50 73.33 1.36 5.27 0.21c
b b b b
No protection 42.50 53.33 1.66 5.36 0.24a
C.V. (%) 3.78 6.54 1.51 4.63 1.36
Average values in each column followed by the same letter are statistically equal (Duncan’s MRT, p [ 0.05). C.V, coefficient of
variation

protection or with the graft union covered by paraffin,
respectively. Furthermore, the plants protected with a
plastic bag had an average of 2.23 shoots, which were
on average 8.17 cm long, compared to the 1.36 and
1.66 shoots of 5.21 and 2.77 cm in length of the
unprotected and paraffin protected plants,
respectively.
Generally, the grafted plants protected with a
plastic bag performed best; all plants with this
protection system attained a successful graft union,
irrespective of the grafting technique (Fig. 3). Con-
versely, the lowest grafting success was observed for
the straight cut graftings protected with liquid paraffin
(17%), but the cleft and splice grafting techniques also
resulted in a low percentage of graft unions formed
when protected with liquid paraffin (25 and 25.5%,
respectively).
Discussion
General discussion
Global agrobiodiversity is decreasing, with fewer crop
species providing a larger proportion of human diets;
more than 50% of the world’s plant-derived calories
are supplied by the three major staple crops rice,
wheat, and maize (Bioversity International 2017). At
the same time, malnourishment is widespread (ca.
16% of the world population), the world population is
projected to increase to more than 9 billion in 2050,
and climate change, land degradation, and competition
for water and land resources is prevalent (Kahane et al.
2013). In order to attain food security within a
sustainable development paradigm it may be benefi-
cial to focus on expanding agrobiodiversity by reduc-
ing focus on the ca. 80 crop species providing 90% of
human energy consumption (Prescott-Allen and Pre-
scott-Allen 1990) and increasing focus on the remain-
der of the ca. 7000 plant species that have been used
for food by humans (Hammer 1998). Cultivation and
incorporation of underutilized crops into human and
livestock diets can infer a number of potential benefits,
including improved food security and nutrition, espe-
cially in developing areas, optimized land use through
utilization of marginal lands, and diversified income
opportunities for small scale farmers. Furthermore,
tapping into the resources available from underutilized
crops provides an increased selection of crop species
for addressing climate change and for including in the

123
Genet Resour Crop Evol
Fig. 3 Percentages of graft unions formed as an effect of the
protection systems and grafting techniques
development of sustainable agricultural systems, e.g.,
through incorporation in agroforestry, intercropping
and crop rotation systems. The crop species currently
recommended to achieve sufficient amounts of the
omega-3 fatty acid alpha-linolenic acid (ALA) are
flaxseed, walnut, and canola (Tur et al. 2012).
However, a number of underutilized crop species
exist that are viable alternatives to these recommen-
dations, and may help increase agrobiodiversity. For
example, chia (Salvia hispanica L.) seed oil has a high
content of omega-3 fatty acids, but cultivation of the
crop was considerably reduced after Spanish colo-
nization in pre-Columbian Mesoamerica (Jamboonsri
et al. 2012). In recent years the consumption of chia
has increased considerably and there is a growing
interest in the study of chia (Oliveira-Alves et al.
2017). Similarly, it is likely that the popularity of P.
volubilis will increase due to the nutritionally bene-
ficial properties of the plant. Nevertheless, the domes-
tication of neglected or underutilized crops poses
certain challenges; typically the amount of intraspeci-
fic diversity is high, the quality of the products varies,
and awareness from consumers, farmers, technology
providers, and research funding bodies is low. Acqui-
sition of germplasm, and distribution of seeds or
vegetative propagation material may be difficult.
Accordingly, studies into genetic diversity, previous
and future domestication approaches, and functional
phytochemistry of promising crops is imperative.
Insights into the propagation, intra- and inter-specific
compatibility, and agronomic properties are highly
relevant, as they may aid in addressing the challenges
surrounding emerging crops.
The current study explores vegetative propagation
of P. volubilis through grafting, a technique which
may aid in addressing the challenges of varying
quality, intraspecific diversity, and accessibility of
propagation material.
Discussion of grafting techniques
The percentages of successful graft unions did not
vary significantly between the three different grafting
techniques. There was, however, a tendency for plants
grafted with the cleft grafting technique to perform
better than the two other grafting techniques. This
could be due to the surface area and placement of the
adjacent tissues in the graft union; when grafting, the
tissues of the rootstock and scion, respectively, are
placed in close unison after the sectioning of the
respective stems. Compatible tissues with meristem-
atic competence are then capable of forming a graft
union. This occurs through the production of callus
from recent cambial derivatives in approximately
equal proportions from the two stems (Esau 1965).
The parenchymatic callus tissue fills the gap between
the rootstock and the scion, where the calluses of the
two stems intermingle. Eventually vascular cambium
forms through the callus, at the point where the
cambiums of the rootstock and scion each meet the
callus, and eventually the cambial cells of the two
stems will merge. The new tissues resulting from this
cambium are in continuity with the phloem and xylem
of the rootstock and scion, thus producing a vascular
connection, and the graft union is complete (Esau
1965).
The cleft grafting technique has a larger wound
surface area than the straight grafting technique, and
therefore a larger area of cambium facing the graft
union, and this may explain the tendency for the cleft
grafting technique to perform best. The cleft and splice
grafting techniques result in a similar cambial surface
area at the graft union and may therefore explain why
the two techniques provide similar results. However,
the cleft technique may help to situate the cambium of
the rootstock and scion more precisely against each
other, since it is not as prone to slip during the
attachment of the scion to the rootstock as the
obliquely cut tissues of the splice technique. Accord-
ingly, although not demonstrated by the results of the
current study, the cleft technique may be superior,
however, the splice technique is more rapid and may
123

therefore be preferred. A better alignment of the
wound tissues of the rootstock and scion increases the
area where the cambium produces callus, which
eventually leads to renewed growth of the scion
(Vázquez et al. 1997). Generally, the cleft technique is
the most commonly used technique for grafting woody
forest species such as pine, eucalyptus and mahogany
(Emhart 1998; Filho et al. 2001). Its lesser popularity
for herbaceous species is probably due to the increased
technical difficulties and—often—lack of necessity.
There was a significant difference in the graft union
success, mortality, number of shoots, length of shoots
and diameter of shoots between the three different
protection systems. For all parameters, the plants
protected by a plastic bag performed best. As noted by
Parkinson et al. (1990) one of the most important
factors for successful grafting is to avoid desiccation
of the graft union. With an increasing concentration
difference between the air humidity and the plant
tissue water potential, plant transpiration also
increases (Zeiger et al. 1987). It is very likely that
the plastic bags has kept an increased air humidity
around the grafted P. volubilis compared to the plants
without plastic bags, and this may have reduced the
transpiration. A reduced degree of desiccation may
therefore explain the higher grafting success for the
plants protected with plastic bags.
Generally, the shade with 20% light transparency
can be assumed to have furthered the development of
successful graft unions in the current study, since it
protects the plants from direct sunlight. Direct sunlight
can cause light stress, through the exposure to excess
light energy, which damages DNA, proteins and other
cellular components, e.g., by production of free
oxygen radicals (Müller-Xing et al. 2014). The harvest
of the scions during the early hours of morning is also
likely to have had a positive influence on the outcome
of the graftings, along with the removal of leaves, as
both these approaches will have reduced the plants’
transpiration. Furthermore, the sterilization of equip-
ment prior to grafting was probably also beneficial in
reducing the risk of fungal infections in the graft
union.
Conclusions and recommendations
The domestication and increased utilization of
neglected and underutilized crops has a number of
123
Genet Resour Crop Evol
potential benefits, including increased agrobiodiver-
sity, improved food security, and diversified opportu-
nities for addressing climate change and developing
sustainable agricultural systems. The development of
a vegetative propagation protocol for P. volubilis can
aid in the domestication of this promising crop, and
may alleviate challenges caused by intraspecific
genetic variation and varying quality of the product.
Vegetative propagation of P. volubilis by grafting
was demonstrated to be possible and a protocol has
been developed. In the current study, a 100% graft
union formation was attained by using the cleft
grafting technique, sealing the wound with a plastic
strip and covering the plant with a plastic bag. The
split grafting technique was, however, also found to be
feasible and may be more rapid.
Vegetative propagation of P. volubilis by grafting
can be used for conserving germplasm to be used in
future breeding approaches, and for rapid multiplica-
tion of superior genotypes to be used commercially.
Vegetative propagation by grafting allows the selec-
tion of disease resistant rootstocks and high yielding
scions, together giving a potentially higher and more
secure yield. This can benefit the local farmers in the
plants native range and may help shift cultivation from
illegal crops. Furthermore, production of P. volubilis
can help combat malnutrition among farmers and in
the local communities.
It will be interesting to study the effects of using
mature rootstocks, as this may decrease the time to
first harvest. Mapping of superior genotypes for use as
rootstocks and scions, respectively, will also be very
beneficial. Furthermore, it might prove fruitful to
attempt grafting of other species of Plukenetia, e.g.,
the large seeded Plukenetia huayllabambana R.
W. Bussmann, C. Téllez et A. Glenn, onto a P.
volubilis rootstock.
Acknowledgements We are grateful to Dennis del Castillo
(PROBOSQUE, Perú) and Luis Arévalo Lopez (IIAP, Perú) for
their practical assistance and help in enabling the project.
Compliance with ethical standards
Conflict of interest The authors declare that they have no
conflict of interest.
Genet Resour Crop Evol
References
Arévalo GG (1996) El cultivo del Sacha Inchi (Plukenetia vol-
ubilis L.) en la Amazonia. programa nacional de investi-
gación en recursos genéticos y biotecnologı́a,
PRONARGEB, Estación experimental El Porvenir, Tara-
poto, Perú
Aspajo GG, Rojas TR, del Castillo AMR, Bourdy G, Deharo E
(2016) Traditional ethnobotanical knowledge of Sacha
Inchi Plukenetia volubilis L. in Northern Peruvian Amazon
(abstract). Libro de resumenes del XV Conabot 2016,
Cusco
Barter G (2016) Vegetative propagation. In: Ingram DS, Vince-
Prue D, Gregory PJ (eds) Science and the garden: the sci-
entific basis of horticultural practice, 3rd edn. Wiley, UK,
pp 149–165
Bioversity International (2017) Mainstreaming agrobiodiversity
in sustainable food systems: scientific foundations for an
agrobiodiversity index. Bioversity International, Rome
Bussmann RW, Téllez C, Glenn A (2009) Plukenetia huayl-
labambana sp. nov. (Euphorbiaceae) from the upper
Amazon of Peru. Nordic J Bot 27:313–315
Bussmann RW, Zambrana NP, Téllez C (2013) Plukenetia
carolis-vegae (Euphorbiaceae)—a new useful species from
Northern Peru. Econ Bot 67:387–392
Cachique D (2006) Estudio de la biologı́a floral y reproductiva
en el cultivo de Sacha Inchi Plukenetia volubilis L. Agró-
nomo, Tarapoto, Perú, Universidad Nacional de San Mar-
tı́n, Tesis Ing, p 70
Cachique D, Rodriguez Á, Ruiz-Solsol H, Vallejos G, Solis R
(2011) Propagación vegetative del SachaInchi (Plukenetia
volubilis L.) mediante enraizamiento de estacas juveniles
en cámaras de subirrigación en la Amazonia Peruana. Folia
Amaz 20:95–100
Chirinos R, Zuloeta G, Pedreschi R, Mignolet E, Larondelle Y,
Campos D (2013) Sacha Inchi (Plukenetia volubilis): a
seed source of polyunsaturated fatty acids, tocopherols,
phytosterols, phenolic compounds and antioxidant capac-
ity. Food Chem 141:1732–1739
Cisneros FH, Paredes D, Arana A, Cisneros-Zevallos L (2014)
Chemical composition, oxidative stability and antioxidant
capacity of oil extracted from roasted seeds of Sacha-Inchi
(Plukenetia volubilis L.). J Agric Food Chem
62:5191–5197
Corazon-Guivin M, Rodrı́guez Á, Cachique D, Chota W, Vás-
quez G, del Castillo D, Renno J-F, Garcı́a-Dávila C (2008)
Diversidad genética en poblaciones naturales de Sacha
Inchi Plukenetia volubilis L. (Euphorbiaceae) en el
departamente de San Martı́n (Perú). Folia Amaz 17:83–90
Corazon-Guivin M, Castro-Ruiz D, Chota-Macuyama W,
Rodrı́guez Á, Cachique D, Manco E, Del-Castillo D,
Renno J-F, Garcı́a-Dávila C (2009) Caracterización
genética de accessiones san martinenses del banco nacional
de germoplasma de Sacha Inchi Plukenetia volubilis L.
(E.E. El Porvenir–INIA). Folia Amaz 18:23–31
Emhart SV (1998) Mejora Genética Forestal Operativa. In:
Ipinza R, Gutierrez B, Emhart V (eds) Artes Gráficas y
Centenario LTDA, Chile, pp. 153–165
Esau K (1965) Plant Anatomy, 2nd edn. Wiley, New York
Fanali C, Dugo L, Cacciola F, Beccaria M, Grasso S, Dachá M,
Dugo P, Mondello L (2011) Chemical characterization of
Sacha Inchi (Plukenetia volubilis L.) oil. J Agric Food
Chem 59:12043–13049
Filho ANK, Hoffmann HA, Tavares RF (2001) Mini-garfagem:
Un novo método para a enxertia do mognosul-americano
(Switenia macrophylla King). Comunicado Técnico 62,
Embrapa Florestas
Gillespie LJ (1993) A synopsis of Neotropical Plukenetia
(Euphorbiaceae) including two new species. Syst Bot
18:575–592
Gillespie LJ (2007) A revision of Paleotropical Plukenetia
(Euphorbiaceae) including two new species from Mada-
gascar. Syst Bot 32:780–802
Guillén MD, Ruiz A, Cabo N, Chirinos R, Pascual G (2003)
Characterization of Sacha Inchi (Plukenetia volubilis L.)
oil by FTIR Spectroscopy and 1H NMR. Comparison with
linseed oil. JAOCS 80:755–762
Gutiérrez L-P, Rosada L-M, Jiménez Á (2011) Chemical com-
position of Sacha Inchi (Plukenetia volubilis L.) seeds and
characteristics of their lipid fraction. Grasas Aceites
62:76–83
Hamaker BR, Valles C, Gilman R, Hardmeier RM, Clark D,
Garcia HH, Gonzales AE, Kohlstad I, Castro M, Valdivia
R, Rodriguez T, Lescano M (1992) Amino acid and fatty
acid profiles of the Inca peanut (Plukenetia volubilis).
Cereal Chem 69:461–463
Hammer K (1998) Agrarbiodiversität und pflanzengenetische
Resourcen. Schriften zu genetischen Resourcen, Band 10,
ZADI, Bonn, Germany, p 98. http://www.genres.de/
fileadmin/SITE_GENRES/downloads/schriftenreihe/Band10_
Gesamt.pdf. Accessed 5 Jan 2017
Hanelt P (ed) (2001) Mansfeld’s encyclopedia of agricultural
and horticultural crops. Springer, New York
Jamboonsri W, Phillips TD, Geneve RL, Cahill JP, Hildebrand
DF (2012) Extending the range of an ancient crop, Salvia
hispanica L.—a new x3 source. Genet Resour Crop Evol
59:171–178
Jáuregui AMM, Ureta CA-O, Castañeda BC, Caparó FL,
Mendoza EB, Lucero LC, Céspedes EM (2013) Estudio
Nutricional de Plukenetia huayllabambana sp. Nov Rev
Soc Quı́m Perú 79:47–56
Kahane R, Hodgkin T, Jaenicke H, Hoogendoorn C, Hermann
M, Keatinge JDH, Hughes JA, Padulosi S, Looney N
(2013) Agrobiodiversity for food security, health and
income. Agron Sustain Dev 33:671–693
Krivankova B, Cepkova PH, Ocelak M, Juton G, Bechyne M,
Lojka B (2012) Preliminary study of diversity of
Plukenetia volubilis based on the morphological and
genetic characteristics. Agric Trop Subtrop 45:140–146
Müller-Xing R, Xing Q, Goodrich J (2014) Footprints of the sun:
memory of UV and light stress in plants. Front Plant Sci
5:474
Oliveira-Alves SC, Vendramini-Costa DB, Cazarin CBB, Júnior
MRM, Ferreira JPB, Silva AB, Prado MA, Bronze MR
(2017) Characterization of phenolic compounds in chia
(Salvia hispanica L.) seeds, fiber flour and oil. Food Chem
232:295–305
Parkinson M, O’Neill CM, Dix PJ (1990) Grafting in vitro. In:
Pollard JW, Walker JM (eds) Methods in molecular
123

biology 6: plant cell and tissue culture. Humana Press, New
York, pp 93–105
Prescott-Allen R, Prescott-Allen C (1990) How many plants
feed the world? Conserv Biol 4:365–374
Rodrigues HS, de Oliveira AB, Lopes MTG, Cruz CD, Chaves
FCM, Bentes JLS (2013) Genetic diversity of Sacha Inchi
accessions detected by AFLP molecular markers. Amaz J
Agric Environ Sci 56:55–60
Ruiz-Solsol H, Mesén F (2010) Efecto del ácidoindolbutı́rico y
de estaquilla en el enraizamiento de Sacha Inchi
(Plukenetia volubilis L.). Agron Costarric 34:259–267
123
Genet Resour Crop Evol
Snedecor GW, Cochran WG (1980) Statistical methods, 7th edn.
Iowa State University Press, Iowa
Tur JA, Bibiloni MM, Sureda A, Pons A (2012) Dietary sources
of omega 3 fatty acids: public health risks and benefits. Br J
Nutr 107:S23–S52
Vázquez C, Orozco A, Rojas M, Sánchez ME, Cervantes V
(1997) La reproducción de las plantas: semillas y meris-
temos, 1st edn. Fondo de cultura económica, México
Zeiger E, Farquhar GD, Cowan IR (1987) Stomatal function.
Stanford University Press, Palo Alto
Zobel B, Talbert J (1988) Técnicas de mejoramiento genético de
árboles forestales. Editorial Limusa, México, p 554